Dastych, H. 2011. Bergtrollus Dzimbowski Gen. N., Sp. N., A

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Entomol. Mitt. Zool. Mus. Hamburg 15 (186): 335-359 Hamburg, 1. Dezember 2011 ISSN 0044-5223

Bergtrollus dzimbowski gen. n., sp. n., a remarkable new tardigrade genus and species from the nival zone of the Lyngen Alps, Norway (Tardigrada: Milnesiidae)

HIERONYMUS DASTYCH

(with 51 gures)

Abstract Bergtrollus dzimbowski gen. n., sp. n., a new genus and species of Tardigrada (the Milnesiidae) from moss in the nival zone of the Lyngen Alps (Norway), is described. This is the second milnesiid genus, apart from Milnesium Doyère, 1840, known to occur in the Northern Hemisphere. The new tardigrade is characterized by a strikingly long and protrusible proboscis (‘snout’) formed by the prolonged mouth region. A similar organ in Eutardigrada has also been reported in Milnesioides exsertum Claxton, 1999 and in the recently discovered Limmenius porcellus Horning et al., 1978 (see Claxton 1999) (both Milnesiidae). These species are known from limited records in New Zealand and Australia. The shape and length of the buccal tube in Bergtrollus gen. n. is intermediate between those of Milnesioides and Limmenius, but the mouth cavity is similar to Milnesium. Diagnostic morphological characters and identi cation key for all four genera of the Milnesiidae are presented and the phylogenetic status of the new taxon within the family is discussed.

Keywords: Tardigrada, taxonomy, Bergtrollus dzimbowski gen. n., sp. n., nival zone, the Lyngen Alps, Norway, the Arctic.

Introduction Milnesium tardigradum Doyère, 1840 is one of the rst described, still valid, worldwide recorded and commonly known tardigrades. The species repre- sents well-separated phyletic branch within Eutardigrada de ned by Schuster et al. (1980) as the order Apochela. The order consists of the sole family Milnesiidae Ramazzotti, 1962 with the three genera, Milnesium Doyère, 1840, Limmenius Horning et al., 1978 and Milnesioides Claxton, 1990. Within Milnesium more than 16 nominal species have been described in recent years (for a check-list see e.g., Guidetti & Bertolani 2005, Degma & Guidetti 2007). Two remaining genera are monospeci c and represented by Limmenius porcellus Horning, Schuster & Grigarick, 1978 and Milnesioides exsertum Claxton, 1999. These species, in particular L. porcellus, are characterized by a very long buccal tube, as compared to Milnesium. Moreover,

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Figs 1-2. The type locality of Bergtrollus dzimbowski gen. n., sp. n.: general (1) and detailed (2) view. In the background E slope of the subsummit of Mt. Steindalstinden.

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the uppermost part of the head and mouth region in M. exsertum form a characteristic, extremely long ‘snout’ (Claxton 1999), a structure not known in other Eutardigrades. The presence of the snout (= proboscis) has also been discovered recently in the originally poorly described Limmenius (see Claxton, l.c.). While the members of Milnesium are common and distributed worldwide, those of Limmenius and Milnesioides have been recorded very rarely and only from the Southern Hemisphere. L. porcellus is known from a few localities in New Zealand (Horning et al. 1978) and from a single specimen from Tasmania (Claxton 1999), while M. exsertum has been reported only from Australia, including Tasmania (Claxton, l.c.).

When recently processing bryophytes from the Lyngen Alps (Northern Norway), several milnesiid tardigrades intermediate in their morphology between Limmenius and Milnesioides were encountered. Analysis of the material brought to light a new and remarkable tardigrade genus and species and also the rst record of the second genus of the family Milnesiidae in the Northern Hemisphere. The paper describes the new tardigrade, compares it with other milnesiid genera and discusses its phylogenetic status within the family. Furthermore, a diagnostic key for these genera is provided.

Material and Methods The new tardigrade were found in a small sample of moss (ca 7-10 cm2) collected in the nival zone of the Lyngen Alps, Norway (for details see “type locality” below). The moss was placed in a paper bag and air-dried. After seven months the sample was soaked in distilled water and kept overnight at the temperature of 0 °C. Tardigrades were extracted by the method described by Dastych (1985), with some living individuals being observed under a stereo-microscope. The animals were xed for SEM examination with hot Bouin’s medium, dehydrated, critical-point-dried and carbon-coated or, for light microscope study, killed with hot water and mounted in a Faure’s medium as permanent microslide preparations. One specimen (holotype) was xed in a mix- ture of methanol and acetic acid (3:1), stained with aceto-lactic orcein (see e.g. Berto- lani 1971) and mounted after examination, like other specimens, in Faure’s medium.

Measurements were taken with PHC. The body length was measured without legs IV, mostly with retracted proboscis (‘snout’). The proboscis was fully extended in only two preserved individuals (one mounted for SEM: Figs 3, 11). However, as these animals represent simplex-stage, their lamellar operculum and peribuccal papillae are torn away (Figs 15-17). Consequently, the proboscis was measured without these structures (slide no. 1982A, Table 1, column “b”: 80.1 m). The length of the organ, when available, was obtained indirectly from the remaining individuals (n = 3, holotype included), i.e. through adding lengths of retracted pseudosegments of the proboscis and that of lamellar operculum. The distance between the (upper) edge of the base of the cephalic papilla and the apex of the mouth region was considered as the length of the snout.

The length of the bucco-pharyngeal and buccal apparatus includes peribuccal lamellae. The term ‘buccal apparatus’ includes mouth cavity, buccal tube and associated structures; the ’bucco-pharyngeal apparatus’ embraces additionally the pharynx. The mouth cavity (= ‘mouth ring’ by Dewel & Clark 1973, in part) was measured with and

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without the lamellae. The (external) diameter of the mouth cavity was taken at the bases of the peribuccal lamellae, immediately below their perpendicular striation. The mouth cavity length (that without lamellae) is the distance between the upper (strongly sclerotized) edge of the stylet sheaths and the level of the bases of peribuccal lamellae, just below their striation (comp. Claxton 1999). The length of the buccal tube was measured from the upper edge of the stylet sheaths up to the posterior edge of the tube, including terminal apophyses. The length of the buccal tube is made up of two units, i.e. that from the upper edge of the stylet sheaths up to the anterior edge of the stylet support insertion (= SSA unit; = pt ss index in Tumanov 2006, based on Pilato 1981) and that from the anterior (fore) edge of the stylet support insertion up to the posterior edge of the tube, including its terminal apophyses (= SSB unit). The (external) diameter of the tube is taken just above the insertion of the stylet supports. Two specimens (slide no. T1985c and T1982d) had on the left and right side of the tube different (aberrant) length of the unit SSB. The unit sides measured 58.3 vs. 63.0 and 61.1 vs. 64.8 m, respectively. Arbitrarily, the higher values of these lengths have been considered for calculations.

The whole length of the primary (= main) branch of the claw was measured from the apex of accessory spines up to the external edge of the arch-like cuticular bar (mbb: Figs 30, 31), which bar is located dorsally and laterally at the nger-like base of the branch. The length of sclerotized unit of the primary branch was measured from the apex of accessory spines up to very proximal termination of its sclerotized com- ponent (= “main branch length” in Tumanov 2006: Fig. 1). The length (height) of the remaining (i.e. basal) part of claw (= ‘basal claw’: Fig. 29: bc) includes lunula (= ‘basal thickening’: e.g. Tumanov 2006) and is the distance between the tip of the secondary branch and the posterior edge of the lunula. An additional measurement of the basal claw excludes the lunula. The width of lunula is a diameter of the structure.

The morphometric indices applied are de ned in Dastych (2006, 2011). For calcu- lation of the hind claw base index (“HBI”) only the sclerotized part of the claw main branch was considered [the length (height) of hind basal claw, lunula excluded x 100 / sclerotized unit of the hind claw main branch]. Furthermore, the following new morphometric indices have been introduced:

1) the snout / body index (“SBI”) i.e., the (percent) ratio between the length of the snout (proboscis) and the body;

2) the mouth cavity index (“MCAVI”) which describes the shape of the mouth cavity [the length of mouth cavity, peribuccal lamellae excluded x 100 / mouth cavity width];

3) the hind claw lunula index (“HLI”) which de ned the size of the hind lunula (the hind claw lunula width x 100 / the basal claw length, lunula included).

Microphotographs were taken with ZEISS ‘Fotomikroskop III’, SEM micrographs with LEO 1525. Figures 19, 23, 24, 34 come from the holotype of the new taxon, the other images represent paratypes.

Figs 3-6. Bergtrollus dzimbowski gen. n., sp. n., animal in simplex-stage. 3. habitus: latero-dorsally, 4-6: apex of the proboscis (‘snout’) in latero-dorsal, latero-ventral and lateral view, respectively (peribucal lamellae and papillae torn out). (Scale bars for Figs 5-6: 10 m. Other explanations in text).

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Figs 7-14. Bergtrollus dzimbowski gen. n., sp. n. 7. head region, proboscis partly retracted: lateral view, 8. head, proboscis retracted: latero-dorsally, 9. mouth region, apex of proboscis: laterally (comp. Fig. 7), 10. mouth region: frontoventrally, 11. proboscis in simplex-stage: laterally, 12. mouth region: ventro-laterally, 13. head region: cuticle, pores at the muscle attachment, 14. cephalic papilla and muscle attachment: dorsally. (Explanations in text).

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The new tardigrade has been compared with Milnesium tardigradum Doyère, 1848 from Norway (Jotunheimen, Fig. 41 and Glomfjell). Moreover, four paratypes of Milne- sioides exsertum [borrowed from the Australian Museum, Sydney: microslides No. KS 41615 (see Fig. 51), KS 41616-617 (Figs 44, 45), KS 41618] and four paratypes of Limmenius porcellus have been examined. The latter came from the National Musem of New Zealand Te Papa Tongarewa, Wellington [two microslides: No. NZ 23 (Fig. 43), NZ 378 (Figs 47, 49, 50)] and from the Bohart Museum, University of California, Davis [two slides, No. NZ 8 (Fig. 46), NZ 221 (Fig. 48)].

Abbreviations used are: ar - artefact bc - basal claw bt – buccal tube cp - cephalic (head) papilla DIC - differential interference contrast ey - eye dot gr - groove HBI - the hind claw base index LM - light microscope lu - lunula mar - muscle attachment region mbb - main branch bar MCAVI - mouth cavity index mc - mouth cavity mcra - mouth cavity range n.a. - not applicable ov – ovocyte pb - primary (main) branch ph - pharynx PHC - phase contrast pl - peribuccal lamella pp – peribuccal papilla pr - proboscis (snout) PT - whole buccal tube indices (Pilato 1981; = WTI indices: Dastych 2006) PUI - posterior tube unit indices qa - quaternary branch r2, r squared - coef cient of determination se - secondary branch SBI - snout / body index SD - standard deviation SEM – scanning electron microscope sh - stylet sheath ss - stylet support SSA - buccal tube anterior unit (the distance between stylet sheaths and stylet support insertion) SSB - buccal tube posterior unit (the distance between stylet support and terminal apophyses) st – stylet tb - transversal bar te - tertiaty branch WTI - whole buccal tube indices (= pt indices, Pilato, 1981) WT SSA - stylet supports Aanterior@ index (= pt ss: Tumanov 2006, based on Pilato 1981) V - coef cient of variability ZMH - Zoologisches Museum Hamburg - (arithmetic) mean.

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Description of new genus Class: Eutardigrada (Richters, 1926) emend. Marcus, 1927 Order: Apochela Schuster et al., 1980 Family: Milnesiidae Ramazzotti, 1962

Bergtrollus gen. n.

DIAGNOSIS: Semiterrestrial, medium- to large-sized milnesiids with a very long, telescopically protrusible tubular proboscis (‘snout’). Mouth cavity short. Buccal tube long, exible, smooth (i.e., without net- /spiral-like strengthening), stylets thin and long. The claw system of Milnesium-type.

TYPE SPECIES: Bergtrollus dzimbowski sp. n. by monotypy.

Included taxa: A single species, Bergtrollus dzimbowski sp. n.

ETYMOLOGY: Bergtrollus = mountain troll. The new genus is named after the mystic creatures, trolls, thought to inhabite Scandinavia (Berg = mountain: German, Norwe- gian).

REMARKS: Bergtrollus gen. n. is the third of four known genera of the Mil- nesiidae with strikingly long, protrusible proboscis (‘snout’), this organ being absent in Milnesium itself. The proboscis is in all these taxa of ca. 10-20 % of their body length [SBI index for the new genus: = 15.08 ± 1.56 (13.8- 17.3) (n = 4), Milnesioides: 9.7 and 15.8, Limmenius: = 18.20 ± 3.92 (13.3- 22.8) % (n = 4)].

The buccal tube length of the new genus is intermediate between that of the genera Milnesioides and Limmenius, as is also the width of the tube (comp. Figs 42-44). The shortest and the widest tube (Fig. 41) in the family have members of Milnesium. In Bergtrollus gen. n. the mouth cavity is short and wide [MCAVI index: = 37.60 ± 6.13 (31.3-50.0) (n = 8)] resembling that of Milnesium [ = 33.5 ± 7.04 (26.9-40.9) (n = 4)] but is different in shape and size from the cavity of the genera reported from the Southern Hemisphere, i.e., Milnesioides and Limmenius. The latter taxa, particularly Limmenius, have the mouth cavity distinctly narrower and longer [MCAVI: = 105.80 ± 16.0 (93.8-129.4) (n = 4) and = 157.60 ± 21.09 (128.7-178.0) (n = 4), respectively; see also Figs 41-44].

Figs 15-19. Bergtrollus dzimbowski gen. n., sp. n. (15-17: animal in simplex-stage): 15. habitus, water-mounted ; ‘empty’ proboscis (arrowhead), mouth appendages torn out: lateral view, 16. habitus (arrow: cephalic papilla): ventrally, 17. proboscis in a late simplex-stage: the mouth region and buccal apparatus retracted (asterisks, respectively) and in a process of formation: dorsal view, 18. head region: strongly retracted proboscis and such anterior part of buccal apparatus: dorsally, 19. greatly retracted, telescopically arranged proboscis (arrows) and bucco-pharyngeal apparatus: dorso- ventrally. (All PHC. Other explanations in text).

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The insertion of the stylet supports in Bergtrollus gen. n. is similar to that in Milnesium and Limmenius and the supports in these genera are inserted at ca. 60 % of the buccal tube length [WTSSA (= pt ss) index in the new genus: 60.6-62.2 (n = 3), Milnesium: = 63.2 ± 3.3 (59.6-67.6) (n = 4), Limmenius: = 59.78 ± 3.73 (55.3-62.9) (n = 4)]. In Milnesioides the supports are inserted much more caudally, at ca. 80 % of the tube length ( = 77.03 ± 1.81 (75.4-79.6) % (n = 4). It should be noted that the insertion of the supports (the index values) is more similar in genera with different type of buccal tube (Milnesium vs. Limmenius) but is distinctly different in genera where the buccal apparatus is more similar, i.e., Milnesium vs. Milnesioides (Figs 41-44).

The claw system of Bergtrollus gen. n. is of the same type as that of other milnesiids, indicating its conservative character within the whole family.

The mouth and head appendages and the type of claws place Bergtrollus dzimbowski gen. n., sp. n. without doubt in the family Milnesiidae. However, the phylogenetic position of this taxon within the family is not clear and at present can be only a subject of speculation.

As inferring from the presence of proboscis and the morphology of its buccal tube, the new genus could be considered as an intermediate form between genera Milnesioides and Limmenius. Consequently, it could be placed as a “missing link” in such a position within prearranged series of the three remaining milnesiid genera, as illustrated but not commented by Claxton (1999: Fig. 10a-c). Milnesioides has a broader but shorter mouth tube than Bergtrollus gen. n. but is somehow similar to the tube of Milnesium. On the other hand, in Limmenius the tube is very long, narrower and much more exible than in the new genus. Moreover, in Limmenius the tube is provided with a net- /spiral-like strengthening. The organ is within the Milnesiidae the most advanced. The tube of the latter genus resembles such independently evolved organ of e.g., the genus Diphascon (the order Parachela, the Diphasconidae).

The mouth cavity in Bergtrollus gen. n. is very short and wide and resembles that in Milnesium. The cavity differs markedly from those present in the both Southern Hemisphere genera, Milnesioides and Limmenius. Because of this, one can also hypothesise that the new taxon originated not within the southern (Gondwanan?) lineage to which Milnesioides and Limmenius might belong, but evolved independently from Milnesium-like ancestors within the northern (Laurasian) region.

Figs 20-23. Bergtrollus dzimbowski gen. n., sp. n. 20. mouth region and anterior part of buccal apparatus in greatly retracted proboscis (comp. Fig. 18): dorsally, 21. devel- oping buccal apparatus, the late simplex-stage: dorsally (comp. Fig. 17), 22. bucco- pharyngeal apparatus, proboscis retracted (arrow): dorsally, 23. mouth region and anterior part of buccal apparatus; proboscis telescopically arranged, retracted (arrows): dorsally. (All PHC. Other explanations in text).

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Figs 24-31. Bergtrollus dzimbowski gen. n., sp. n. 24. fragment of buccal apparatus: dorso-ventrally, 25. claws IV, 26. fragment of the cuticle above legs IV: dorsally, 27. claws IV (arrowhead: main branch cuticular bar, mbb): laterally, 28. claws I: laterally, 29. claw II, 30. claws III, 31. basal claws IV. (All PHC. Other explanations in text).

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Apart from scarce information on Milnesium, there is no molecular data concerning other milnesiid genera. Data on their morphology, biology and distribution are very rare. Further studies might thus help to understand bet- ter evolutionary processes within the family Milnesiidae and phyletic relation within its genera.

The diagnostic key for the Milnesiidae genera is provided after the species description.

DISTRIBUTION: Known only from the type locality in the Lyngen Alps, North- ern Norway (the Arctic zone).

Description of the species Bergtrollus dzimbowski sp. n. (Figs 3-40, 42)

TYPE MATERIAL. Holotype. – (Figs 19-23, 24, 34). Female, 666 m long, coll. H. Dastych, 13 Sept 2009. Stained with lactic orcein, mounted in Faure’s medium. The microslide (No. T1985j, slide No. 1) is deposited in the Zoologisches Museum Hamburg (ZMH Acc. No. A28/11).

Type locality. – (Figs 1-2). Northern Norway, Troms, the Lyngen Alps. Upper region of the glacier Steindalsbreen. A small pass between Steindal- and Goverdal Valley, at SW ridge of the Nállangáisi-Massiv (N 69° 23’ 48.6’’+ E 019° 49’ 52.2’’, ca. 1000 m a.s.l.). A scree eld on the pass at the base of a small semi-nunatak S nxen. The rock moss Andreaea rupestris Hedw. (Andreaeceae) on a gabbro (silicate) boulder, not shaded, exposed SE; 13 Sept. 2009, coll. H. Dastych.

Paratypes. – Locality data as above. Altogether 10 females. Eight animals mounted in Faure’s medium on ve slides (Nos 2-6: ZMH Acc. No. A29/11). One para- type is deposited at the Zoological Museum Bergen, Norway (slide No. 4), the Aus- tralian Museum, Sydney (slide No. 2), and the National Museum of New Zealand Te Papa Tongarewa, Wellington (slide No. 3). The remainder of type specimens is lodged in the ZMH, including two paratypic individuals mounted on SEM stubs (ZMH Acc. No. A30/11).

The species co-occurred in the sample with Echiniscus wendti Richters, 1903, Hebesuncus conjungens (Thulin, 1911), Macrobiotus cf. islandicus Richters, 1904, Milnesium eurystomum Maucci, 1991 and Mil. cf. tardigradum Doyére, 1840.

ETYMOLOGY. – The speci c name is in honour of my fellow climber and friend, Hans- Jochen Dzimbowski (Deutscher Alpenverein, Hamburg), the organizer of our trips to the Norwegian mountains.

DIAGNOSIS. – As for genus.

DESCRIPTION. – Body 492-736 (holotype 666) m long, light-brown or yel- low-brownish, with darker pigmented areas in its rear. Cuticle smooth (Fig. 13), sometimes in small areas slightly folded (Fig. 26). Eye-dots relatively large (Figs 18, 32, 33), composed of irregularly shaped and differently sized pigment granules.

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Mouth region located on a long, protrusible tubular proboscis (‘snout’) (Figs 3, 11, 16-18: pr; Fig. 15: arrowhead), more or less retracted in preserved specimens (e.g., Figs 7, 8, 10, 19). A small but distinct vertical ring- like groove around anterior part of the proboscis, at its ca. 1/3 distal length (Figs 4-6, 11, 17, 18, 20: gr). The groove divides the proboscis externally in two differently long pseudosegments (Fig. 17) when the ‘snout’ is fully extended or retracted. The tubular proboscis retracts mostly in the form of a usually telescopically closely arranged, two or three shorter subunits (Figs 19, 22, 23, 33: arrows). When fully extended, it is ca. 15 % of the body length (SBI = 13.8-17.3 %, n = 4). Two cephalic papillae symmetrically are located at the basal part of the proboscis (Figs 7, 10: cp).

Bucco-pharyngeal apparatus large, almost half of the body length. Mouth cavity small, buccal tube markedly long, exible (Figs 19, 20, 21, 32, 33: bt). Pharynx large, pear-shaped (Figs 15, 19: ph).

Figs 32-36. Bergtrollus dzimbowski gen. n., sp. n. 32-33: buccal apparatus, 34. basal claw IV, 35. claws I, 36. basal claws IV. (All DIC. Other explanations in text).

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Mouth opening surrounded by six nger-like peribuccal papillae (Figs 8, 9, 18, 20: pp) and covered by an operculum composed of six peribuccal lamellae. The lamellae triangular, elongated (8-10, 23: pl), with a ne vertical striation, visible in LM (Fig. 20) and partly in SEM. Mouth cavity small, short, its lining strongly sclerotized (Figs 18, 20, 23, 32: mc), its wall smooth, without discernible armature (teeth or granules). The openings of stylet sheaths leading into the mouth cavity small, their location strongly sclerotized (Figs 18, 20, 23). The posterior part of sheaths weakly sclerotized, shaped as a long and delicate cuticular tubes (Figs 20, 23, 32: sh). Just below the strongly sclerotized (anterior) units of the stylet sheaths two small, roundish, differently light-defracting elds. These areas are lighter and less sclerotized than the surrounding wall of the buccal tube (Fig. 20: arrowhead, 42). The structures represent a part of the pharyngeal organs (see note in § “Re- marks”).

Buccal tube very long, relatively wide and exible (Figs 19, 22, 33: bt). The tube wall smooth, i.e. without net- /spiral-like strengthening. The tube slightly thickened at its posterior end, thus forming small and at terminal apophyses (Figs 32, 33: arrowhead). As in other milnesiids, the tube is located (disembogues) just at the uppermost anterior edge of the pharynx. No placoids in the pharynx lumen. Stylets long, markedly thin (Figs 22, 32, 33: st), with a broad, at spade-like furca (Fig. 24: fu). Stylet supports located in ca. 62 % of the tube length (Figs 22, 33: ss). One specimen (microslide No. 6) in the simplex-stage was in a process of development of its buccal apparatus, which was thus weakly sclerotized. The buccal tube remainded there ‘soft’, its wall being distinctly folded (Fig. 21).

Claws of Milnesium-type. Each claw composed of an apical unit, i.e. the primary (= main) branch (Figs 30, 39: pb) and well separated remaining part, the basal unit (Fig. 29: bc). The primary branch short, slightly thickened, with distinct accessory spines (Figs 28, 40). The branch base dorsally and laterally with an arch-like cuticular bar (Figs 30, 31, 34: mbb). The basal unit of the claw (= basal claw: Fig. 29: bc) composed of three branches, i.e. the secondary, tertiary and the lowest, quaternary branch (Figs 30, 39: se, te, qa, respectively). The latter branch has also been termed as ‘basal spur’: e.g., Tumanov 2006). The branches relatively long, their number (3) on particular claw constant. The proximal part of the basal claw (i.e., its foot) with a roundish structure, the lunula (Figs 31, 40: lu), particularly well formed on legs IV. (The latter called also ‘basal thickening’: e.g., Pilato & Binda 2010, Tumanov l.c.). Below basal claws I-III occurs ventrally a transversal cuticular bar (Figs 29, 30: tb). Claws I not modi ed, indicating thus all animals examined as females.

Eggs unknown.

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Figs 37-40. Bergtrollus dzimbowski gen. n., sp. n. 37. claws I, 38. claws II, 39. claws IV, 40. claws IV. (Explanations in text).

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Morphometric data Measurements are in m, all indices in %. Their values are presented as follows:

± SD (min-max) [n] * V (for measurements); ± SD (min-max) [n] * V / r2 (for indices).

For the abbreviations and de nitions of indices see § “Material and methods” and Dastych (2006, 2011). The morphometrics of the holotype (666 m long) is separated from other data by a dot (•) and marked in bold. Not measured (unfavourably posi- tioned) structures in the holotype are marked with A /- A.

A) Measurements (m)

a. Body length 592.70 ± 78.78 (492.8-736.4) [9] * 13.3 • 666.4 b. Proboscis (‘snout’) length 95.98 ± (80.1-106.7) [4] * 13.1 • 91.8 c. Peribuccal papilla length n.a. (5.4, 6.3) [2] * n.a. • / - d. Cephalic papilla length 6.48 ± 0.75 (5.4-7.2) [5] * 11.6 • 5.4 e. Bucco-pharyngeal apparatus length 278.20 ± 36.87 (231.0-330.0) [5] * 13.3 • 330.0

f. Pharynx length 108.90 ± 11.11 (90.9-123.2) [8] * 10.2 • 123.2 g. Pharynx width 55.86 ± 10.48 (45.0-77.0) [8] *18.8 • 77.0 h. Mouth cavity length (+lamellae) 13.28 ± 1.25 (10.8-14.4) [8] * 9.42 • 14.4 i. Mouth cavity length (- lamellae) 4.85 ± 1.03 (4.1-7.2) [8] * 21.3 • 4.5 j. Mouth cavity (ext.) width 12.94 ± 1.86 (10.8-16.2) [8] * 14.4 • 14.4 k. Stylet sheaths length 27.00 ± 2.55 (24.3-29.7) [5] * 9.4 • 29.7 l. Buccal tube length n.a. (153.0-171.9) [3] * n.a. • 171.9 m. SSA length (tube above stylet supports) n.a. (92.7-103.5) [3] * n.a. • 103.5 n. SSB length (tube below stylet supports) 62.74 ± 6.05 (49.5-67.5) [7] * 9.7 • 65.7 o. Buccal tube (ext.) width 8.35 ± 1.37 (6.8-9.9) [8] * 16.4 • 9.9

p. Ext. claw 1 whole main branch length 20.80 ± 2.93 (16.2-22.5) [9] * 9.3 • 22.5 q. Ext. claw 1 sclerot. main branch length 17.50 ± 1.80 (13.5-19.8) [9] * 10.3 • 19.8 r. Ext. claw 1 basal unit (+ lunula) length 14.60 ± 1.54 (11.7-16.2) [9] * 10.6 • 16.2 s. Ext. claw 1 basal unit (- lunula) length 12.50 ± 1.49 (9.0-13.1) [8] * 12.0 • 13.5 t. Ext. claw 1 lunula width 3.32 ± 0.41 (2.7-3.6) [8] * 12.3 • 3.6

u. Hind claw whole main branch length 22.95 ± 1.55 (21.6-25.2) [4] * 6.8 • / - v. Hind claw sclerot. main branch length 19.44 ± 0.49 (18.9-19.8) [5] * 2.5 • 18.9 w. Hind claw basal unit (+ lunula) length 15.63 ± 1.72 (12.6-18.0) [7] * 11.2 • 18.0 x. Hind claw basal unit (- lunula) length 13.28 ± 1.50 (10.8-15.3) [6] * 11.3 • 15.3 y. Hind claw lunula width 5.11 ± 0.36 (4.5-5.4) [6] * 7.0 • 5.4

For a list of measurements taken from each type specimen see Appendix 1.

B) Indices

1) WTI (the whole tube length indices) (= “pt indices”: Pilato 1981).

WT proboscis (‘snout’) length n.a. (53.4-69.7) [3] * n.a. / 84.0 • 53.4 WT cephalic papilla length n.a. (3.1, 4.1) [2] * n.a. / - • 3.1 WT buccal apparatus length n.a. (180.6-192.0) [3] * n.a. / 95.7 • 192.0 WT pharynx length n.a. (71.6-74.1) [3] * n.a. / 91.7 • 71.6 WT pharynx width n.a. (32.8-44.8) [3] * n.a. / 24.7 • 44.8

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WT mouth cavity length (+lamel.) n.a. (8.4-9.4) [3] * n.a. / - • 8.4 WT mouth cavity length (- lamel.) n.a. (2.6-4.5) [3] * n.a. / 12.5 • 2.6 WT mouth cavity (ext.) width n.a. (8.4-10.6 [3] * n.a. / 72.6 • 8.4 WT stylet sheaths length n.a. (17.3-18.8 [3] * n.a. / 27.4 • 17.3 WT SSA length (tube above stylet supports) n.a. (60.6-62.2) [3] * n.a. / 90.8 • 60.2 WT SSB length (tube below stylet supports) n.a. (38.2-41.2) [3] * n.a. / 77.3 • 38.2 WT buccal tube (ext.) width n.a. (5.8-6.5) [3] * n.a. / < 0.1 • 5.8

WT ext. claw 1 whole main branch length n.a. (13.1-14.7) [3] * n.a. / < 0.1 • 13.1 WT ext. claw 1 sclerot. main branch length n.a. (11.1-11.8) [3] * n.a. / 77.3 • 11.5 WT ext. claw 1 basal unit (+ lunula) length n.a. (9.4-10.6) [3] * n.a. / < 0.1 • 9.4 WT ext. claw 1 basal unit (- lunula) length n.a. (7.9-8.8) [3] * n.a. / - • 7.9 WT ext. claw 1 lunula width n.a. (2.1-2-2) [3] * n.a. / 72.6 • 2.1

WT hind claw whole main branch length n.a. (13.9, 16.5) [2] * n.a. / - • - WT hind claw sclerot. main branch length n.a.. (11.0-12.9) [3] * n.a. / 72.6 • 11.0 WT hind claw basal unit (+ lunula) length n.a. (10.0-10.6) [3] * n.a. / 77.3 • 10.5 WT hind claw basal unit (- lunula) length n.a. (8.6-8.9) [3] * n.a. / 95.1• 8.9 WT hind claw lunula width n.a. (3.1-3.5) [3] * n.a. / < 0.1 • 3.1

2) PUI (the posterior tube unit indices)

PU proboscis (‘snout’) length 159.50 ± 13.56 (139.7-169.4) [4] * 8.5 / 54.5 • 139.7 PU peribuccal papilla length n.a. (8.5-9.6) [2] * n.a. / n.a. • - PU cephalic papilla length 10.60 ± 1.72 (8.2-12.7) [5] * 16.2 / 0.2 • 8.2 PU buccal apparatus length 452.20 ± 42.18 (401.9-502.3) [4] * 9.3 / 20.5 • 502.3

PU pharynx length 174.5 ± 14.30 (155.6-191.4) [6] * 8.2 / 0.5 • 187.5 PU pharynx width 88.45 ± 18.53 (69.4-117.2) [6] * 21.0 /1.8 • 117.2 PU mouth cavity length (+lamel.) 21.05 ± 1.72 (19.2-22.9) [6] * 8.2 / 19.1 • 21.9 PU mouth cavity length (- lamel.) 7.77 ± 1.95 (6.3-11.4) [6] * 25.1 / 48.5 • 6.8 PU mouth cavity (ext.) width 20.60 ± 3.55 (16.4-25.7) [6] * 17.2 / 53.6 • 21.9 PU stylet sheaths length 45.43 ± 6.86 (37.5-54.2) [4] * 15.1 / 0.4 • 54.2 PU buccal tube (ext.) width 13.57 ± 2.11 (11.0-15.7) [6] * 15.8 /8.8 • 15.1

PU ext. claw 1 whole main branch length 33.00 ± 2.10 (29.3-35.7) [7] * 6.4 / 62.3 • 34.2 PU ext. claw 1 sclerot. main branch length 27.49 ± 1.91 (24.7-30.1) [7] * 7.0 / 59.5 • 30.1 PU ext. claw 1 basal unit (+ lunula) length 23.21 ± 2.28 (18.7-25.7) [7] * 9.8 / 31.6 • 24.7 PU ext. claw 1 basal unit (- lunula) length 20.10 ± 1.22 (18.4-21.4) [6] * 6.1 / 81.4 • 20.5 PU ext. claw 1 lunula width 5.48 ± 0.20 (5.1-5.7) [6] * 3.7 / 87.4 • 5.5 PU hind claw whole main branch length n.a. (32.9-40.0) [3] * n.a. / n.a. • - PU hind claw sclerot. main branch length 30.08 ± 1.06 (28.8-31.4) [4] * 3.5 /- • 28.8 PU hind claw basal unit (+ lunula) length 24.87 ± 2.12 (21.3-27.4) [6] * 8.5 / 46.1 • 27.4 PU hind claw basal unit (- lunula) length 21.62 ± 1.43 (19.4-23.3) [5] * 6.6 / 70.2 • 23.3 PU hind claw lunula width 8.42 0.46 (7.9-9.1) [5] * 5.4 / 86.2 • 8.2

Figs 41-47. Types of the buccal apparatus in the Milnesiidae. 41. Milnesium tardigradum Doyère, 42. Bergtrollus dzimbowski gen. n., sp. n., 43. Limmenius porcellus Horn- ing et al., 44-45: Milnesioides exsertum Claxton, buccal apparatus and its anterior part, respectively, 46-47: L. porcellus, anterior part of buccal apparatus: dorso-ventral and lateral view, respectively. (All PHC. The range of mouth cavity (mcra) is marked by arrows. Paratypes used for Figs 42-47 are indicated in § “Material and methods”. Other explanations in text).

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C) Other indices

Snout / body index (SBI) 15.03 ± 1.56 (13.8-17.3) [4] * 10.4 / 45.5 • 13.8 Mouth cavity index (MCAVI) 37.60 ± 6.13 (31.3-50.0) [8] * 16.3 / 31.4 • 31.3 Hind claw base index (HBI) 71.88 ± 6.14 (68.2-81.0) [4] * 8.6 / 61.0 • 81.0 Hind claw lunula index (HLI) 29.40 ± 2.72 (29.4-35.7) [6] * 8.4 / 54.8 • 30.0

VARIABILITY. – The individuals of B. dzimbowski sp. n. examined are characterized by a relatively low intra-speci c variability of their qualitative fea- tures. As the morphometric data of the new species are based only on few available measurements (n = 2-9), the results of descriptive statistics have often only an approximate character. The coef cient of determination of the WT SSA (= pt ss) index is high, with r squared value equalling 90.8 % (n = 3). It indicates a high degree of correlation between the length of the buccal tube and SSA unit. The index is considered as of high diagnostic importance (Pilato 1981) and already standardized for eutardigrades. In the new species it ranges between 60.6-62.2 % (n = 3).

REPRODUCTION MODE AND PLOIDY. – Only females have been found among all (11) specimens of B. dzimbowski sp. n. examined. It is therefore too early to try to de ne the reproduction mode in the species. In all genera

Figs 48-51. Limmenius porcellus Horning et al. (48-50): 48. cuticular pattern at the body rear: dorso-laterally, 49. net- /spiral-like structure of the buccal tube, 50. anterior part of the head region, proboscis (arrows) and buccal apparatus partly retracted: dorso-ventrally, 51. Milnesioides exsertum Claxton, cuticular pattern at the body rear: dorso-laterally. (All PHC. Paratypes used for Figs 48-51 are indicated in § “Material and methods”. Other explanations in text).

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of the Milnesiidae the bisexual mode prevails and occurs at least in several species of Milnesium. This mode has also been reported for Milnesioides exsertum (see Claxton 1999). In the present study three males were found among examined paratypes of Limmenius porcellus (slides Nos. NZ 8, 23, 378), indicating bisexual reproduction in this genus also (unpublished). The ploidy in B. dzimbowski sp. n. is not known.

BIOLOGY AND DISTRIBUTION. – The limited biological information on the new species can only be partly inferred from data on its type locality in the Lyngen Alps (Figs 1, 2). B. dzimbowski sp. n. has been found in a nival zone of the Norwegian continental Arctic, in a small patch of rock moss, Andrea- cea rupestris Hedw. on a gabbro boulder. According to Nyholm (1981), the moss inhabits siliceous rocks and occurs from the woodland up to the high- alpine belt of the mountains, being common in the north and central part of Fennoscandia.

Live individuals of B. dzimbowski sp. n. move forward in the same way as described already for Milnesioides exsertum by Claxton (1999: 187), i.e., “… the extended “snout” was observed moving from side to side as if animal was searching for food…”. The animals easily retract their proboscis when disturbed, then greatly resembling members of Milnesium. The type of diet of the new species is unknown, as in the intestines of examined specimens no plant material, rotifer or nematode remnants could be observed.

REMARKS. – The phylogenetic position of Bergtrollus dzimbowski sp. n. within the Milnesiidae has been discussed at the description of new genus.

The pharyngeal organs of B. dzimbowski sp. n. represent a chemosensory structural complex discussed already in tardigrades e.g., by Dewel & Clark 1973, Dewel & Eibye-Jacobsen 2006, Walz 1978, Greven 1980, Wiederhöft & Greven 1999. A part of the organ, two lighter, roundish elds within the mouth tube wall (e.g., Fig. 20: arrowheads) correspond supposedly to the perforated areas of the tube (comp. Greven 1980: Fig. 17). Such perforation have been already described by the authors (l.c.) in Macrobiotus and Mil- nesium and is discernible only with the electron microscope. Within the Mil- nesiidae these organs, apart from Milnesium and Bergtrollus gen. n., occur also in Milnesioides (see Figs 44, 45: unpublished) and have been observed as small hardly detectable elds in two of four examined paratypes of Lim- menius porcellus (unpublished).

The term ‘lunula’ applied here for the roundish structure at the base (foot) of the basal claw in milnesiids does not indicate a homology with similarly located structures in remote phyletic lineages of Eutardigrada, such as in most macrobiotids or many hypsibiids s.l. The origin of lunulae and their phyletic relations in all these taxa is unclear. Supposedly the structures evolved independently as some other structures did within eutardigrades. Among observed genera of the Milnesiidae with a proboscis, the lunulae are relatively the largest in B. dzimbowski sp. n.

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Key to the genera of the family Milnesiidae

1. Buccal tube short, wide (Fig. 41; the values of the WT buccal tube width index ca. 40 %), no proboscis (= snout) ...... Milnesium

– Buccal tube long or very long, narrower (Figs 41-43; the WT buccal tube width index lower than 20 %), proboscis* present (Figs 15, 50) ...... 2

2. Stylet supports inserted caudally (Fig. 44; the WTSSA index ca. 80 %), buccal tube relatively wide (the WT buccal tube width index ca. 20 %) ...... Milnesioides

– Stylet supports inserted more medially (Figs 41-43; WTSSA ca. 60%), buccal tube distinctly longer, narrower (the WT buccal tube width index lower than 10 %) ...... 3

3. Mouth cavity short, wide (Figs 20, 42; the MCAVI index ca. 40 %, n = 3), buccal tube smooth (no net- /spiral-like strengthening), shorter and more rigid ...... Bergtrollus gen. n.

– Mouth cavity much longer, narrower (Fig. 43; MCAVI ca. 160 %, n = 4), buccal tube with net- /spiral-like strengthening (Fig. 49) ** . . . Limmenius

Zusammenfassung Eine neue Bärtierchengattung und -art, Bergtrollus dzimbowski gen. n., sp. n. (Tar- digrada) aus Moos der nivalen Zone der Lyngen Alpen in Nordnorwegen wird be- schrieben. Charakteristisch für die neue Art ist der außerordentlich verlängerte und herausziehbare Teil der Mundregion (= Proboscis). Bei Eutardigrada wurde ein sol- ches Organ nur in den Gattungen Milnesioides Claxton, 1999 und Limmenius Hor- ning et al., 1978 festgestellt, die nur selten von Neuseeland und Australien gemeldet wurden. Bergtrollus dzimbowski gen. n., sp. n. steht durch die Form und Länge der Mundröhre zwischen den Gattungen Milnesioides und Limmenius. Durch die Ähnlich- keit der Mundhöhle steht sie nah an der Gattung Milnesium. Diagnostische Merkmale von vier bekannten Gattungen der Familie Milnesiidae und der phylogenetische Status innerhalb der Familie werden diskutiert. Ein Bestimmungsschlüssel für diese Gattun- gen wurde erstellt.

* The milnesiid individuals with strongly retracted proboscis (see e.g. Figs 19, 23) resemble super cially members of Milnesium under low (stereo-) microscope mag- ni cation. ** The net- /spiral-like strengthening of the mouth tube in Limmenius is weakly de- veloped (see Claxton 1999: Fig. 9), often discernible only when its picture is strongly contrasted in an image processing application (Fig. 49).

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Acknowledgements I am very grateful to Hans-Jochen Dzimbowski (DAV, Hamburg), an enthusiast and expert of Norway, for providing the opportunity to collect this spectacular animal. I am much obliged to Dr. Ricardo Palma (National Museum of New Zealand Te Papa Tongarewa, Wellington, Dr. Steven L. Heydon (the Bohart Museum, University of Cali- fornia, Davis) and Dr. Graham Milledge (the Australian Museum, Sydney) who kindly loaned me the type specimens from the collections in their care. I thank very much Prof. Dr. Georg Gärtner (Botanisches Institut, Universität Innsbruck) for identi cation of moss sample, his valuable remarks and reference data, Prof. Dr. Hartmut Greven (Institut für Zoologie, Universität Düseldorf) for his very helpful comments and Renate Walter for her assistance in obtaining SEM micrographs. I am thankful to Dr. Dietrich L. Bürkel (Hamburg) for the English linguistic revision. All support for this project from the University Hamburg is gratefully acknowledged.

References Bertolani, R. 1971: Contributo alla cariologia dei Tardigradi. Osservazioni su Macrobiotus hufelandii. – Atti Acc. Naz. Lincei, Rend., 50 (6): 772-775. Roma.

Claxton, S.K. 1999: Milnesioides exsertum gen. n. sp. n., a new tardigrade from Australia (Tardigrada: Milnesiidae). – Zool. Anz., 238: 183-190. Jena.

Dastych, H. 1985: West Spitsbergen Tardigrada. – Acta zool. cracov., 28 (3): 169-214. Krakow.

Dastych, H. 2006: A new tardigrade species of the genus Ramazzottius Binda & Pilato, 1986 (Tardigrada) from the nival zone of the Mont Blanc Massive (The Western Alps), with some morphometric remarks. – Mitt. hamb. Zool. Mus. Inst., 103: 33-45. Hamburg.

Dastych, H. 2011: Ramazzottius agannae sp. nov., a new tardigrade species from the nival zone of the Austrian Central Alps (Tardigrada). – Entomol. Mitt. Zool. Mus. Hamburg, 15: 237-253. Hamburg.

Degma, P. & Guidetti, R. 2007: Notes to the current checklist of Tardigrada. – Zootaxa, 1579: 41-53. Auckland.

Dewel, R.A. & Clark, W.H. 1973: Studies on the tardigrades. I. Fine structure of the anterior foregut of Milnesium tardigradum. – Tissue & Cell, 5 (1): 133-146. Churchill Livingstone.

Dewel, R.A. & Eibye-Jacobsen, J. 2006: The mouth cone and mouth ring of Echiniscus viridissimus Peter , 1956 (Heterotardigrada) with comparisons to corresponding structures in other tardigrades. – Hydrobiologia, 558: 41-51. Dodrecht.

Greven, H. 1980: Die Bärtierchen – Die Neue Brehm Bücherei, A. Ziemsen Verlag, 101 pp., Wittenberg Lutherstadt.

Guidetti, R. & Bertolani, R. 2005: Tardigrade taxonomy: an updated check list of the taxa and a list of characters for their identi cation. – Zootaxa, 845: 1-46. Auckland.

Horning, D.S., Schuster, R.O. & Grigarick, A.A.: Tardigrada of New Zealand. – NZeal. J. Zool., 5: 185-280. Wellington.

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Nyholm, E. 1981: Illustrated moos ora of Fennoscandia, II. Musci. – Bot. Soc. Lund., 2nd ed., 799 pp. Lund.

Pilato, G. 1981: Analisi di nuovi caratteri nello studio degli Eutardigradi. – Animalia, 8 (1/3): 51-57. Catania.

Pilato, G. & Binda, M.G. 2010: De nition of families, subfamilies, genera and subgenera of the Tardigrada, and keys to their identi cation. – Zootaxa, 2404: 1-54. Auckland.

Schuster, R.O., Nelson, D.R., Grigarick, A.A. & Christenberry, D. 1980: Systematic criteria of the Eutardigrada. – Trans. Amer. Microsc. Soc., 99 (3): 284-303. Washington.

Tumanov, D.V. 2006: Five new species of the genus Milnesium (Tardigrada, Eutardigrada, Milnesiidae). – Zootaxa, 1122: 1-23. Auckland.

Walz, B. 1978: Electron microscopic investigation of cephalic sense organs of the tardigrade Macrobiotus hufelandi C.A.S. Schultze. – Zoomorphology, 89: 1-19. Berlin.

Wiederhöft, H. & Greven, H. 1999: Notes on head sensory organs of Milnesium tardigradum Doyère, 1840 (Apochela, Eutardigrada). – Zool. Anz., 238: 338-346. Jena.

Address of the author:

Dr. H. Dastych, Biozentrum Grindel und Zoologisches Museum, Universität Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany (e-mail: [email protected] hamburg.de).

14674 - Entomologische Mitteilungen - Band 15 - Nummer 186.pdf 54 13.12.2011 09:05:40 Bergtrollus dzimbowski gen. n., sp. n. 359 ) bold lighted ( gen. n., sp. n. [the abbreviations (a-y) 49.5 16.2 13.5 11.7 9.0 2.7 12.6 10.8 4.5 Bergtrollus dzimbowski . Measurements (m) of diagnostic characters in the type series abcdefghijklmnopqrstuvwxy 666.4 91.8 5.4 330.0 123.2 77.0 14.4 4.5 14.4 29.7 171.9 103.5 65.7 9.9 22.5 19.8 16.2 13.5 3.6 18.9 18.0 15.3 5.4 No. No. Slide 1985-J 1985-I 512.21985-H 736.41985-G 105.3 616.01985-C 106.7 492.81982-A 7.2 543.2 292.6 120.6 80.1 53.1 277.2 14.4 113.4 7.2 6.3 65.7 14.4 115.2 14.4 49.5 27.9 231.0 5.4 162.0 13.5 16.2 100.8 90.9 4.5 28.8 63.0 48.6 11.7 153.0 9.6 10.8 21.6 92.7 4.5 18.0 63.0 11.7 15.3 9.9 13.5 22.5 3.6 18.0 22.5 16.2 18.9 13.5 16.2 3.2 67.5 14.0 25.2 9.0 5.0 19.8 19.8 16.2 17.1 13.5 12.6 5.4 6.8 21.6 18.0 14.4 12.6 2.7 14.4 1982-B 537.61982-C 608.41982-D 621.6 6.3 7.2 5.4 6.3 106.2 260.4 54.0 100.8 12.6 45.0 4.5 12.6 100.8 10.8 4.1 54.0 12.6 13.5 24.3 4.1 11.7 24.3 64.8 65.7 7.2 7.2 21.6 20.7 18.0 16.2 14.4 15.3 12.2 7.2 12.6 3.6 20.7 3.6 18.9 21.6 15.3 19.8 13.1 3.6 22.5 15.3 19.8 12.6 16.7 5.4 13.5 5.0 Appendix 1 measurements are those of the holotype]. for particular character are explained in § “Morphometric data”, i.e., ‘a’ = body length, ‘b’ = proboscis length, etc. The high = proboscis length, etc. = body length, ‘b’ for particular character are explained in § “Morphometric data”, i.e., ‘a’

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